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JERUSALEM — Amid the horrors of the Holocaust, the atrocities perpetrated by a few brutal women have always stood out, like aberrations of nature.

There were notorious camp guards like Ilse Koch and Irma Grese. And lesser-known killers like Erna Petri, the wife of an SS officer and a mother who was convicted of shooting to death six Jewish children in Nazi-occupied Poland, or Johanna Altvater Zelle, a German secretary accused of child murder in the Volodymyr-Volynskyy ghetto in Nazi-occupied Ukraine.

The Nazi killing machine was undoubtedly a male-dominated affair. But according to new research, the participation of German women in the genocide, as perpetrators, accomplices or passive witnesses, was far greater than previously thought.

The researcher, Wendy Lower, an American historian now living in Munich, has drawn attention to the number of seemingly ordinary German women who willingly went out to the Nazi-occupied eastern territories as part of the war effort, to areas where genocide was openly occurring.

“Thousands would be a conservative estimate,” Lower said in an interview in Jerusalem last week.

While most did not bloody their own hands, the acts of those who did seemed all the more perverse because they operated outside the concentration-camp system, on their own initiative.

Lower’s findings shed new light on the Holocaust from a gender perspective, according to experts, and have further underlined the importance of the role of the lower echelons in the Nazi killing apparatus.

Lower began traveling to Ukraine in the early 1990s, as the Soviet archives opened up. She started in Zhytomyr, about 75 miles west of Kiev, where the SS leader Heinrich Himmler had his Ukrainian headquarters, and where she found original German files, some burned at the edges, in the local archive.

She noticed the frequency with which women were mentioned at the scenes of genocide.

Women also kept cropping up as witnesses in West and East German investigations after the war.

There were up to 5,000 female guards in the concentration camps, making up about 10 percent of the personnel.

Grese was hanged at the age of 21 for war crimes committed in Auschwitz and Bergen-Belsen; Koch was convicted of participating in murders at Buchenwald.

Only 1 or 2 percent of the perpetrators were women, according to Lower. But in many cases where genocide was taking place, German women were very close by.

Several witnesses have described festive banquets near mass-shooting sites in the Ukrainian forests, with German women providing refreshment for the shooting squads whose work often went on for days.

Petri was married to an SS officer who ran an agricultural estate in occupied Poland. She later confessed to having murdered six Jewish children, aged 6 to 12. She came across them while out riding in her carriage. She was the mother of two young children, and was 25 at the time.

Near naked, the Jewish children had apparently escaped from a railroad car bound for the Sobibor camp. She took them home, fed them, then led them into the woods and shot them one by one.

She told her interrogators that she had done so, in part, because she wanted to prove herself to the men.

She was tried in East Germany and served a life sentence.

Altvater Zelle went to Ukraine as a 22-year-old single woman and became the secretary of a district commissar, Wilhelm Westerheide. Survivors remembered her as the notorious Fraulein Hanna, and accused her, among other things, of smashing a toddler’s head against a ghetto wall and of throwing children to their deaths from the window of a makeshift hospital.

One survivor, Moses Messer, said he saw the woman he knew as Fraulein Hanna smashing the toddler to death against the wall. He told lawyers in Haifa, Israel, in the early 1960s: “Such sadism from a woman I have never seen. I will never forget this scene.”

Lower met the daughter of Petri, who died in 2000, in a tiny hamlet in the former East Germany.